Sensory-specific Areas Research Articles

The Relevance of Ecological Transitions to Intelligence in Marine Mammals.

Macphail’s comparative approach to intelligence focused on associative processes, an orientation inconsistent with more multifaceted lay and scientific understandings of the term. His ultimate emphasis on associative processes indicated few differences in intelligence among vertebrates. We explore options more attuned to common definitions by considering intelligence in terms of richness of representations of the world, the interconnectivity of those representations, the ability to flexibly change those connections, and knowledge. We focus on marine mammals, represented by the amphibious pinnipeds and the aquatic cetaceans and sirenians, as animals that transitioned from a terrestrial existence to an aquatic one, experiencing major changes in ecological pressures. They adapted with morphological transformations related to streamlining the body, physiological changes in respiration and thermoregulation, and sensory/perceptual changes, including echolocation capabilities and diminished olfaction in many cetaceans, both in-air and underwater visual focus, and enhanced senses of touch in pinnipeds and sirenians. Having a terrestrial foundation on which aquatic capacities were overlaid likely affected their cognitive abilities, especially as a new reliance on sound and touch, and the need to surface to breath changed their interactions with the world. Vocal and behavioral observational learning capabilities in the wild and in laboratory experiments suggest versatility in group coordination. Empirical reports on aspects of intelligent behavior like problem-solving, spatial learning, and concept learning by various species of cetaceans and pinnipeds suggest rich cognitive abilities. The high energy demands of the brain suggest that brain-intelligence relationships might be fruitful areas for study when specific hypotheses are considered, e.g., brain mapping indicates hypertrophy of specific sensory areas in marine mammals. Modern neuroimaging techniques provide ways to study neural connectivity, and the patterns of connections between sensory, motor, and other cortical regions provide a biological framework for exploring how animals represent and flexibly use information in navigating and learning about their environment. At this stage of marine mammal research, it would still be prudent to follow Macphail’s caution that it is premature to make strong comparative statements without more empirical evidence, but an approach that includes learning more about how animals flexibly link information across multiple representations could be a productive way of comparing species by allowing them to use their specific strengths within comparative tasks.

Lower Beta: A Central Coordinator of Temporal Prediction in Multimodal Speech.

How the brain decomposes and integrates information in multimodal speech perception is linked to oscillatory dynamics. However, how speech takes advantage of redundancy between different sensory modalities, and how this translates into specific oscillatory patterns remains unclear. We address the role of lower beta activity (~20 Hz), generally associated with motor functions, as an amodal central coordinator that receives bottom-up delta-theta copies from specific sensory areas and generate top-down temporal predictions for auditory entrainment. Dissociating temporal prediction from entrainment may explain how and why visual input benefits speech processing rather than adding cognitive load in multimodal speech perception. On the one hand, body movements convey prosodic and syllabic features at delta and theta rates (i.e., 1–3 Hz and 4–7 Hz). On the other hand, the natural precedence of visual input before auditory onsets may prepare the brain to anticipate and facilitate the integration of auditory delta-theta copies of the prosodic-syllabic structure. Here, we identify three fundamental criteria based on recent evidence and hypotheses, which support the notion that lower motor beta frequency may play a central and generic role in temporal prediction during speech perception. First, beta activity must respond to rhythmic stimulation across modalities. Second, beta power must respond to biological motion and speech-related movements conveying temporal information in multimodal speech processing. Third, temporal prediction may recruit a communication loop between motor and primary auditory cortices (PACs) via delta-to-beta cross-frequency coupling. We discuss evidence related to each criterion and extend these concepts to a beta-motivated framework of multimodal speech processing.

Sex Differences in the Olfactory System: a Functional MRI Study

Olfactory dysfunction is an early marker of neurological disease and a common symptom in psychotic disorders. Previous anatomical and functional research suggests that sex effects may be crucial in the assessment of the olfactory system. Nonetheless, the neural mechanisms through which the factor sex impacts olfactory perception are still not well understood. In this context, we use fMRI to investigate sex differences in the passive processing of chemical stimuli, in order to obtain new neuroscientific data that may help improve the assessment of odor perception. Thirty healthy subjects (17 women) were stimulated with mint and butanol (event-related design) in a 3.0-T MRI scanner. A one-sample t test analysis was performed in order to observe olfactory-related activations. Intergroup differences (women vs. men) and the influence of each aroma were analyzed using a 2 × 2 ANOVA and post hoc contrasts. Men and women showed differential activity (males > females) in right superior/middle temporal areas, the right inferior frontal cortex, and the hypothalamus. Both groups showed a predominance of the right hemisphere for the processing of odors. Functional differences between women and men in olfaction are not restricted to specific sensory areas and reflect a more general sex-dependent effect in multisensory integration processes. Considering sex differences is essential in order to develop more specific and efficient strategies for the assessment and rehabilitation of the olfactory system and for the interpretation of the olfactory loss as an early biomarker of neurological and psychiatric diseases.

Occipital alpha power reveals fast attentional inhibition of incongruent distractors.

Recent associative models of cognitive control hypothesize that cognitive control can be learned (optimized) for task-specific settings via associations between perceptual, motor, and control representations, and, once learned, control can be implemented rapidly. Midfrontal brain areas signal the need for control, and control is subsequently implemented by biasing sensory representations, boosting or suppressing activity in brain areas processing task-relevant or task-irrelevant information. To assess the timescale of this process, we employed EEG. In order to pinpoint control implementation in specific sensory areas, we used a flanker task with incongruent flankers shown in only one hemifield (congruent flankers in the other hemifield) isolating their processing in the contralateral hemisphere. ERPs revealed fast modulations specifically in visual processing areas contralateral to the incongruent flankers. To test whether these modulations reflect increased or decreased processing of incongruent flankers, we investigated alpha power, a marker for attentional inhibition. Importantly, we show increased alpha power over visual areas processing incongruent flankers from 300 to 500 ms poststimulus onset. This suggests fast cognitive control by attentional inhibition for information disrupting goal-oriented actions. Additionally, we show that midfrontal theta earlier in the trial is also modulated by incongruency, and that theta power predicts subsequent alpha power modulations. This supports the hypothesis that midfrontal incongruency detection leads to control implementation, and reveals that these mechanisms take place on a fast, within-trial timescale.

Oral texture influences the neural processing of ortho- and retronasal odors in humans

Eating implies mutual interactions between different senses. In the present work we aimed at studying relations between food texture and food odor, using both psychophysical and imaging techniques.Eighteen right-handed healthy human subjects participated to both behavioral and fMRI sessions. Fresh, sweetened milk and a more thickened version were delivered orally; in addition, a buttery-cream aroma was presented ortho- or retronasally. Stimuli were applied using a gustometer and or an air-dilution olfactometer, both computer-controlled. In each session subjects rated separately odor-, taste- and thickness intensities of the stimuli.The behavioral data show that odors, presented through either retro- or orthonasal path, induce a significant flavor enhancement with respect to the no-odor condition. Brain functional data indicated a significant enhancement of the activation of olfactory eloquent areas in favor of ortho-nasal odor presentation while activations of mechanosensory areas were favored by the retro-nasal odor route. As effect of oral stimuli we found a significant correlation between the texture intensity rating vs. the BOLD signal in the supplementary motor area, known to drive subconsciously primed movement, putatively associated in this case with the tongue movement required with the handling of the stimulus. Moreover, we found inhibition of the signal in different sensory specific areas as an effect of the mutual interaction between stimulus qualities.In conclusion, ortho- and retronasal odors differentially affect the neural processing of the texture of oral stimuli.

Neuromodulation of Early Multisensory Interactions in the Visual Cortex

Merging information derived from different sensory channels allows the brain to amplify minimal signals to reduce their ambiguity, thereby improving the ability of orienting to, detecting, and identifying environmental events. Although multisensory interactions have been mostly ascribed to the activity of higher-order heteromodal areas, multisensory convergence may arise even in primary sensory-specific areas located very early along the cortical processing stream. In three experiments, we investigated early multisensory interactions in lower-level visual areas, by using a novel approach, based on the coupling of behavioral stimulation with two noninvasive brain stimulation techniques, namely, TMS and transcranial direct current stimulation (tDCS). First, we showed that redundant multisensory stimuli can increase visual cortical excitability, as measured by means of phosphene induction by occipital TMS; such physiological enhancement is followed by a behavioral facilitation through the amplification of signal intensity in sensory-specific visual areas. The more sensory inputs are combined (i.e., trimodal vs. bimodal stimuli), the greater are the benefits on phosphene perception. Second, neuroelectrical activity changes induced by tDCS in the temporal and in the parietal cortices, but not in the occipital cortex, can further boost the multisensory enhancement of visual cortical excitability, by increasing the auditory and tactile inputs from temporal and parietal regions, respectively, to lower-level visual areas.

Disinhibited feedback as a cause of synesthesia: Evidence from a functional connectivity study on auditory-visual synesthetes

In synesthesia, certain stimuli to one sensory modality lead to sensory perception in another unstimulated modality. In addition to other models, a two-stage model is discussed to explain this phenomenon, which combines two previously formulated hypotheses regarding synesthesia: direct cross-activation and hyperbinding. The direct cross-activation model postulates that direct connections between sensory-specific areas are responsible for co-activation and synesthetic perception. The hyperbinding hypothesis suggests that the inducing stimulus and the synesthetic sensation are coupled by a sensory nexus area, which may be located in the parietal cortex. This latter hypothesis is compatible with the disinhibited feedback model, which suggests unusual feedback from multimodal convergence areas as the cause of synesthesia. In this study, the relevance of these models was tested in a group (n=14) of auditory-visual synesthetes by performing a functional connectivity analysis on functional magnetic resonance imaging (fMRI) data. Different simple and complex sounds were used as stimuli, and functionally defined seed areas in the bilateral auditory cortex (AC) and the left inferior parietal cortex (IPC) were used for the connectivity calculations. We found no differences in the connectivity of the AC and the visual areas between synesthetes and controls. The main finding of the study was stronger connectivity of the left IPC with the left primary auditory and right primary visual cortex in the group of auditory-visual synesthetes. The results support the model of disinhibited feedback as a cause of synesthetic perception but do not suggest direct cross-activation.

Bypassing Primary Sensory Cortices—A Direct Thalamocortical Pathway for Transmitting Salient Sensory Information

Detection and appropriate reaction to sudden and intense events happening in the sensory environment is crucial for survival. By combining Bayesian model selection with dynamic causal modeling of functional magnetic resonance imaging data, a novel analysis approach that allows inferring the causality between neural activities in different brain areas, we demonstrate that salient sensory information reaches the multimodal cortical areas responsible for its detection directly from the thalamus, without being first processed in primary and secondary sensory-specific areas. This direct thalamocortical transmission of multimodal salient information is parallel to the processing of finer stimulus attributes, which are transmitted in a modality-specific fashion from the thalamus to the relevant primary sensory areas. Such direct thalamocortical connections bypassing primary sensory cortices provide a fast and efficient way for transmitting information from subcortical structures to multimodal cortical areas, to allow the early detection of salient events and, thereby, trigger immediate and appropriate behavior.

Noradrenergic and cholinergic modulation of olfactory bulb sensory processing

Neuromodulation in sensory perception serves important functions such as regulation of signal to noise ratio, attention, and modulation of learning and memory. Neuromodulators in specific sensory areas often have highly similar cellular, but distinct behavioral effects. To address this issue, we here review the function and role of two neuromodulators, acetylcholine (Ach) and noradrenaline (NE) for olfactory sensory processing in the adult main olfactory bulb. We first describe specific bulbar sensory computations, review cellular effects of each modulator and then address their specific roles in bulbar sensory processing. We finally put these data in a behavioral and computational perspective.

Tuning of the Human Neocortex to the Temporal Dynamics of Attended Events

Previous studies raise the hypothesis that attentional bias in the phase of neocortical excitability fluctuations (oscillations) represents a fundamental mechanism for tuning the brain to the temporal dynamics of task-relevant event patterns. To evaluate this hypothesis, we recorded intracranial electrocortical activity in human epilepsy patients while they performed an audiovisual stream selection task. Consistent with our hypothesis, (1) attentional modulation of oscillatory entrainment operates in a distinct network of areas including auditory, visual, posterior parietal, inferior motor, inferior frontal and superior midline frontal cortex, (2) the degree of oscillatory entrainment depends on the predictability of the stimulus stream, and (3) the attentional phase shift of entrained oscillation cooccurs with classical attentional effects observed on phase-locked evoked activity in sensory-specific areas but seems to operate on entrained low-frequency oscillations that cannot be explained by sensory activity evoked at the rate of stimulation. Thus, attentional entrainment appears to tune a network of brain areas to the temporal dynamics of behaviorally relevant event streams, contributing to its perceptual and behavioral selection.

The Sensory Representation of Time

The Sensory Representation of Time

Motor Learning with Augmented Feedback: Modality-Dependent Behavioral and Neural Consequences

Sensory information is critical to correct performance errors online during the execution of complex tasks and can be complemented by augmented feedback (FB). Here, 2 groups of participants acquired a new bimanual coordination pattern under different augmented FB conditions: 1) visual input reflecting coordination between the 2 hands and 2) auditory pacing integrating the timing of both hands into a single temporal structure. Behavioral findings revealed that the visual group became dependent on this augmented FB for performance, whereas the auditory group performed equally well with or without augmented FB by the end of practice. Functional magnetic resonance imaging (fMRI) results corroborated these behavioral findings: the visual group showed neural activity increases in sensory-specific areas during practice, supporting increased reliance on augmented FB. Conversely, the auditory group showed a neural activity decrease, specifically in areas associated with cognitive/sensory monitoring of motor task performance, supporting the development of a control mode that was less reliant on augmented FB sources. Finally, some remnants of brain activity in sensory-specific areas in the absence of augmented FB were found for the visual group only, illustrating ongoing reliance on these areas. These findings provide the first neural account for the "guidance hypothesis of information FB," extensively supported by behavioral research.

Audiovisual Temporal Correspondence Modulates Human Multisensory Superior Temporal Sulcus Plus Primary Sensory Cortices

The brain should integrate related but not unrelated information from different senses. Temporal patterning of inputs to different modalities may provide critical information about whether those inputs are related or not. We studied effects of temporal correspondence between auditory and visual streams on human brain activity with functional magnetic resonance imaging (fMRI). Streams of visual flashes with irregularly jittered, arrhythmic timing could appear on right or left, with or without a stream of auditory tones that coincided perfectly when present (highly unlikely by chance), were noncoincident with vision (different erratic, arrhythmic pattern with same temporal statistics), or an auditory stream appeared alone. fMRI revealed blood oxygenation level-dependent (BOLD) increases in multisensory superior temporal sulcus (mSTS), contralateral to a visual stream when coincident with an auditory stream, and BOLD decreases for noncoincidence relative to unisensory baselines. Contralateral primary visual cortex and auditory cortex were also affected by audiovisual temporal correspondence or noncorrespondence, as confirmed in individuals. Connectivity analyses indicated enhanced influence from mSTS on primary sensory areas, rather than vice versa, during audiovisual correspondence. Temporal correspondence between auditory and visual streams affects a network of both multisensory (mSTS) and sensory-specific areas in humans, including even primary visual and auditory cortex, with stronger responses for corresponding and thus related audiovisual inputs.

Multisensory stimulation with or without saccades: fMRI evidence for crossmodal effects on sensory-specific cortices that reflect multisensory location-congruence rather than task-relevance

During covert attention to peripheral visual targets, presenting a concurrent tactile stimulus at the same location as a visual target can boost neural responses to it, even in sensory-specific occipital areas. Here, we examined any such crossmodal spatial-congruence effects in the context of overt spatial orienting, when saccadic eye-movements were directed to each peripheral target or central fixation maintained. In addition, we tested whether crossmodal spatial-congruence effects depend on the task-relevance of visual or tactile stimuli. On each trial, subjects received spatially congruent (same location) or incongruent (opposite hemifields) visuo-tactile stimulation. In different blocks, they made saccades either to the location of each visual stimulus, or to the location of each tactile stimulus; or else passively received the multisensory stimulation. Activity in visual extrastriate areas and in somatosensory parietal operculum was modulated by spatial congruence of the multisensory stimulation, with stronger activations when concurrent visual and tactile stimuli were both delivered at the same contralateral location. Critically, lateral occipital cortex and parietal operculum showed such crossmodal spatial effects irrespective of which modality was task relevant; and also of whether the stimuli were used to guide eye-movements or were just passively received. These results reveal crossmodal spatial-congruence effects upon visual and somatosensory sensory-specific areas that are relatively ‘automatic’, determined by the spatial relation of multisensory input rather than by its task-relevance.

Hodological characterization of the septum in anuran amphibians: II. Efferent connections

The efferent connections of the septum of the gray treefrog Hyla versicolor were studied by combining anterograde and retrograde tracing with biotin ethylendiamine (Neurobiotin). The lateral septal complex projects mainly to the medial pallium, limbic regions (e.g., amygdala and nucleus accumbens), and hypothalamic areas but also to sensory nuclei in the diencephalon and midbrain. The central septal complex strongly innervates the medial pallium, limbic, and hypothalamic areas but also specific sensory (including olfactory) regions. The medial septal complex sends major projections to all olfactory nuclei and a weaker projection to the hypothalamus. Our results indicate that all septal nuclei may modify the animal's internal state via efferents to limbic and hypothalamic areas. Via projections to the medial pallium, lateral and central septal complexes may be involved in learning processes as well. Because of their connections to specific sensory areas, all septal areas are in a position to influence sensory processing. Furthermore, our data suggest that both the postolfactory eminence and the bed nucleus of the pallial commissure are not part of the septal complex, rather, the postolfactory eminence seems to be comparable to the mammalian primary olfactory cortex, whereas the bed nucleus may be analogous to the mammalian subfornical organ.

Dynamics of cortico-subcortical cross-modal operations involved in audio-visual object detection in humans.

Very recently, a number of neuroimaging studies in humans have begun to investigate the question of how the brain integrates information from different sensory modalities to form unified percepts. Already, intermodal neural processing appears to depend on the modalities of inputs or the nature (speech/non-speech) of information to be combined. Yet, the variety of paradigms, stimuli and technics used make it difficult to understand the relationships between the factors operating at the perceptual level and the underlying physiological processes. In a previous experiment, we used event-related potentials to describe the spatio-temporal organization of audio-visual interactions during a bimodal object recognition task. Here we examined the network of cross-modal interactions involved in simple detection of the same objects. The objects were defined either by unimodal auditory or visual features alone, or by the combination of the two features. As expected, subjects detected bimodal stimuli more rapidly than either unimodal stimuli. Combined analysis of potentials, scalp current densities and dipole modeling revealed several interaction patterns within the first 200 micro s post-stimulus: in occipito-parietal visual areas (45-85 micro s), in deep brain structures, possibly the superior colliculus (105-140 micro s), and in right temporo-frontal regions (170-185 micro s). These interactions differed from those found during object identification in sensory-specific areas and possibly in the superior colliculus, indicating that the neural operations governing multisensory integration depend crucially on the nature of the perceptual processes involved.

Generators of the late cognitive potentials in auditory and visual oddball tasks

Recordings directly within the brain can establish local evoked potential generation without the ambiguities always associated with extracranial electromagnetic measures. Depth recordings have found that sensory stimuli activate primary cortex and then material-specific encoders. Sensory-specific areas remain active for long periods, but by about 200 ms are joined by activation in widespread brain systems. One system is related to the orientation of attention. It is centered in paralimbic and attentional frontoparietocingular cortex, and associated with the P3a. A second system associated with P3b envelopes cognitive contextual integration. It engages the ventral temporofrontal event-encoding cortices (inferotemporal, perirhinal, and ventrolateral prefrontal), association cortices (superior temporal sulcal and posterior parietal), and the hippocampus. Thus, even in simple tasks, activation is widespread but concentrated in particular multilobar systems. With this information, the late cognitive potentials can be used to monitor the probable location, timing and intensity of brain activation during cognitive tasks.

Cortex, spikes and waves: the silent areas revisited

The concept of attractor neural networks (ANN) has led to biologically plausible models for neocortical working-associative memory, depending on excitatory feedback within modules of 103-105 neurons, asynchronous updating by units within discrete intervals approximating the spike refractory period and the stabilisation of attractors as different modular patterns of correlated burst firing which would persist for many seconds (of Hebbian reverberatory circuits). Recent studies of up to 11 units simultaneously, in nonspecific cortex of behaving monkeys, confirm traditional impressions of relatively low firing rates and apparently little bursting correlation in these conventionally ‘silent areas’ (in contrast to sensorimotor-specific neocortex) during a bimodal-sensory, complex localisation task. Considering the supposed cognitive functions of nonspecific cortex, the intracortical pathways involved, and not least the data from population-vector and oscillatory slow-wave studies in sensory-specific areas of t...

Inter- and intra-modal processing of sensory-specific stimuli.

Inter- and intra-modal processing was compared with modality-specific forms in two experiments on a group of 40 university students. In Exp. I, the RTs for tactual-to-visual and visual-to-visual processing were compared for 20 subjects, and in Exp. II, the visual-to-tactual and the tactual-to-tactual processing were compared for another group of 20 subjects. Analysis showed that intra-modal processing is faster than inter-modal processing. Further, the "same" RT is faster than the "different" RT. The inter- and intra-modal processing differences are based on the delay in retrieval and recognition across sensory-specific areas.

POSTERIOR DIVISION OF THE EIGHTH CRANIAL NERVE IN LACERTA VIVIPARA.

LITTLE information is available in the literature on the innervation of sensory areas in the inferior, or saccular, portion of the inner ear of the lizard. In order adequately to show nervous structure, histological techniques have been used in the past that preclude dealing with the intact animal; usually the brain must be removed, thereby disrupting anatomical relationships. A notable exception is the work of Beccari1 on embryonic material of Lacerta ; his primary interest, however, was in finding homologies in the central nervous system between reptiles and mammals, and he gives little consideration to the central terminations of fibres from specific sensory areas. Weston2,3 has speculated on central terminations of sensory fibres in Sphenodon and has discussed structure of vestibular and auditory areas in brains of a number of reptiles.